Echolocation transmitting beam of the Atlantic bottlenose dolphin

The transmitting beam patterns of echolocation signals emitted by an Atlantic bottlenose dolphin Tursiops truncatus were measured in the vertical and horizontal planes with an array of seven hydrophones. Particular emphasis was placed on accurately verifying the animal's position on a bite-plate/tail-rest stationing device using underwater video monitoring equipment. The major axis of the vertical beam was directed at an angle of 5 degrees above the plane defined by the animal's lips. This angle was 15 degrees lower than previously measured. The vertical beam measurements indicate that the major axis of the transmitting beam is aligned with the major axis of the receiving beam. The horizontal beam was directed forward. The directivity index of 26.5 dB calculated from the beam pattern measured in both planes agreed well with previous calculation of 25.4 dB.     

Echolocation in the Risso's dolphin,Grampus griseus

The Risso's dolphin (Grampus griseus) is an exclusively cephalopod-consuming delphinid with a distinctive vertical indentation along its forehead. To investigate whether or not the species echolocates, a female Risso's dolphin was trained to discriminate an aluminum cylinder from a nylon sphere (experiment 1) or an aluminum sphere (experiment 2) while wearing eyecups and free swimming in an open-water pen in Kaneohe Bay, Hawaii. The dolphin completed the task with little difficulty despite being blindfolded. Clicks emitted by the dolphin were acquired at average amplitudes of 192.6 dB re 1 microPa, with estimated sources levels up to 216 dB re 1 microPa-1 m. Clicks were acquired with peak frequencies as high as 104.7 kHz (Mf(p) = 47.9 kHz), center frequencies as high as 85.7 kHz (Mf(0) = 56.5 kHz), 3-dB bandwidths up to 94.1 kHz (M(BW) = 39.7 kHz), and root-mean-square bandwidths up to 32.8 kHz (M(RMS) = 23.3 kHz). Click durations were between 40 and 70 micros. The data establish that the Risso's dolphin echolocates, and that, aside from slightly lower amplitudes and frequencies, the clicks emitted by the dolphin were similar to those emitted by other echolocating odontocetes. The particular acoustic and behavioral findings in the study are discussed with respect to the possible direction of the sonar transmission beam of the species.     

Representing multiple discrimination cues in a computational model of the bottlenose dolphin auditory system

A computational model of the dolphin auditory system was developed to describe how multiple discrimination cues may be represented and employed during echolocation discrimination tasks. The model consisted of a bank of gammatone filters followed by half-wave rectification and low pass filtering. The output of the model resembles a spectrogram; however, the model reflects temporal and spectral resolving properties of the dolphin auditory system. Model outputs were organized to represent discrimination cues related to spectral, temporal and intensity information. Two empirical experiments, a phase discrimination experiment [Johnson et al., Animal Sonar Processes and Performance (Plenum, New York, 1988)] and a cylinder wall thickness discrimination tasks [Au and Pawolski, J. Comp. Physiol. A 170, 41-47 (1992)] were then simulated. Model performance was compared to dolphin performance. Although multiple discrimination cues were potentially available to the dolphin, simulation results suggest temporal information was used in the former experiment and spectral information in the latter. This model's representation of sound provides a more accurate approximation to what the dolphin may be hearing compared to conventional spectrograms, time-amplitude, or spectral representations.     

High-frequency auditory filter shape for the Atlantic bottlenose dolphin

High-frequency auditory filter shapes of an Atlantic bottlenose dolphin (Tursiops truncatus) were measured using a notched noise masking source centered on pure tone signals at frequencies of 40, 60, 80 and 100?kHz. A dolphin was trained to swim into a hoop station facing the noise/signal transducer located at a distance of 2?m. The dolphin's masked threshold was determined using an up-down staircase method as the width of the notched noise was randomly varied from 0, 0.2, 04, 0.6, and 0.8 times the test tone frequency. The masked threshold decreased as the width of the notched increased and less noise fell within the auditory filter associated with the test tone. The auditory filter shapes were approximated by fitting a roex (p,r(r)) function to the masked threshold results. A constant-Q value of 8.4 modeled the results within the frequency range of 40 to 100 kHz relatively well. However, between 60 and 100?kHz, the 3?dB bandwidth was relatively similar between 9.5 and 10?kHz, indicating a constant-bandwidth system in this frequency range The mean equivalent rectangular bandwidth calculated from the filter shape was approximately 16.0%, 17.0%, 13.6% and 11.3% of the tone frequencies of 40, 60, 80, and 100?kHz.     

Echolocation signals of wild Atlantic spotted dolphin (Stenella frontalis)

An array of four hydrophones arranged in a symmetrical star configuration was used to measure the echolocation signals of the Atlantic spotted dolphin (Stenella frontalis) in the Bahamas. The spacing between the center hydrophone and the other hydrophones was 45.7 cm. A video camera was attached to the array and a video tape recorder was time synchronized with the computer used to digitize the acoustic signals. The echolocation signals had bi-modal frequency spectra with a low-frequency peak between 40 and 50 kHz and a high-frequency peak between 110 and 130 kHz. The low-frequency peak was dominant when the signal the source level was low and the high-frequency peak dominated when the source level was high. Peak-to-peak source levels as high as 210 dB re 1 microPa were measured. The source level varied in amplitude approximately as a function of the one-way transmission loss for signals traveling from the animals to the array. The characteristics of the signals were similar to those of captive Tursiops truncatus, Delphinapterus leucas and Pseudorca crassidens measured in open waters under controlled conditions.     

Echolocation range of captive and free-ranging baiji (Lipotes vexillifer), finless porpoise (Neophocaena phocaenoides), and bottlenose dolphin (Tursiops truncatus).

The interclick intervals of captive dolphins are known to be longer than the two-way transit time between the dolphin and a target. In the present study, the interclick intervals of free-ranging baiji, finless porpoises, and bottlenose dolphins in the wild and in captivity were compared. The click intervals in open waters ranged up to 100-200 ms, whereas the click intervals in captivity were in the order of 4-28 ms. Echolocation of free-ranging dolphins appears to adapt to various distance in navigation or ranging, sometimes up to 140 m. Additionally, the difference of waveform characteristics of clicks between species was recognized in the frequency of maximum energy and the click duration.     

Comparison of target detection capabilities of the beluga and bottlenose dolphin

The echolocation detection capabilities of a beluga (Delphinapterus leucas) and an Atlantic bottlenose dolphin (Tursiops truncatus) were directly compared in a target detection experiment. Both animals were trained to detect targets in the presence of masking noise. Targets were stainless-steel, water-filled spheres 7.62 and 22.86 cm in diameter. Target ranges of 16.5 and 40 m were used with the 7.62-cm sphere and 80 m with the 22.86-cm sphere. Masking noise with a flat spectrum from 40-160 kHz was projected from a spherical transducer placed 4 or 5 m, depending on the target distance, from the animal hoop station in line with the target. Target detection performance was determined as a function of masking noise level at each target range. The echo-to-noise ratio (Ee/No)max for the beluga at the 75% correct response threshold was approximately 1.0 dB compared to about 10 dB for the dolphin. The differences of each animal's detection performance across the three ranges were consistent with target strength and transmission loss differences. It is speculated that the difference in performance between the two species may be due to differences in critical bandwidth, signal processing capability, or echolocation strategy.     

Acoustic features of objects matched by an echolocating bottlenose dolphin

The focus of this study was to investigate how dolphins use acoustic features in returning echolocation signals to discriminate among objects. An echolocating dolphin performed a match-to-sample task with objects that varied in size, shape, material, and texture. After the task was completed, the features of the object echoes were measured (e.g., target strength, peak frequency). The dolphin's error patterns were examined in conjunction with the between-object variation in acoustic features to identify the acoustic features that the dolphin used to discriminate among the objects. The present study explored two hypotheses regarding the way dolphins use acoustic information in echoes: (1) use of a single feature, or (2) use of a linear combination of multiple features. The results suggested that dolphins do not use a single feature across all object sets or a linear combination of six echo features. Five features appeared to be important to the dolphin on four or more sets: the echo spectrum shape, the pattern of changes in target strength and number of highlights as a function of object orientation, and peak and center frequency. These data suggest that dolphins use multiple features and integrate information across echoes from a range of object orientations.     

Discrimination of complex synthetic echoes by an echolocating bottlenose dolphin

Bottlenose dolphins (Tursiops truncatus) detect and discriminate underwater objects by interrogating the environment with their native echolocation capabilities. Study of dolphins' ability to detect complex (multihighlight) signals in noise suggest echolocation object detection using an approximate 265-micros energy integration time window sensitive to the echo region of highest energy or containing the highlight with highest energy. Backscatter from many real objects contains multiple highlights, distributed over multiple integration windows and with varying amplitude relationships. This study used synthetic echoes with complex highlight structures to test whether high-amplitude initial highlights would interfere with discrimination of low-amplitude trailing highlights. A dolphin was trained to discriminate two-highlight synthetic echoes using differences in the center frequencies of the second highlights. The energy ratio (delta dB) and the timing relationship (delta T) between the first and second highlights were manipulated. An iso-sensitivity function was derived using a factorial design testing delta dB at -10, -15, -20, and -25 dB and delta T at 10, 20, 40, and 80 micros. The results suggest that the animal processed multiple echo highlights as separable analyzable features in the discrimination task, perhaps perceived through differences in spectral rippling across the duration of the echoes.     

Synthesis and modification of the whistles of the bottlenose dolphin, Tursiops truncatus

A signal-processing algorithm was developed to analyze harmonic frequency-modulated sounds, to modify the parameters of the analyzed signal, and to synthesize a new analytically specified signal that resembles the original signal in specified features. This algorithm was used with dolphin whistles, a frequency-modulated harmonic signal that has typically been described in terms of its contour, or pattern of modulation of the fundamental frequency. In order to test whether other features may also be salient to dolphins, the whistle analysis calculates the energies at the harmonics as well as the fundamental frequency of the whistle. The modification part of the algorithm can set all of these energies to a constant, can shift the whistle frequency, and can expand or compress the time base or the frequency of the whistle. The synthesis part of the algorithm then synthesizes a waveform based upon the energies and frequencies of the fundamental and first two harmonics. These synthetic whistles will be useful for evaluating what acoustic features dolphins use in discriminating different whistles.     

Monaural and binaural hearing directivity in the bottlenose dolphin: evoked-potential study

Hearing thresholds as a function of sound-source azimuth were measured in bottlenose dolphins using an auditory evoked potential (AEP) technique. AEP recording from a region next to the ear allowed recording monaural responses. Thus, a monaural directivity diagram (a threshold-vs-azimuth function) was obtained. For comparison, binaural AEP components were recorded from the vertex to get standard binaural directivity diagrams. Both monaural and binaural diagrams were obtained at frequencies ranging from 8 to 128 kHz in quarter-octave steps. At all frequencies, the monaural diagram demonstrated asymmetry manifesting itself as: (1) lower thresholds at the ipsilateral azimuth as compared to the symmetrical contralateral azimuth and (2) ipsilateral shift of the lowest-threshold point. The directivity index increased with frequency: at the ipsilateral side it rose from 4.7 to 17.8 dB from 11.2 to 128 kHz, and from 10.5 to 15.6 dB at the contralateral side. The lowest-threshold azimuth shifted from 0 degrees at 90-128 kHz to 22.5 degrees at 8-11.2 kHz. The frequency-dependent variation of the lowest-threshold azimuth indicates the presence of two sound-receiving apertures at each head side: a high-frequency aperture with the axis directed frontally, and a low-frequency aperture with the axis directed laterally.     

Discrimination of amplitude-modulated synthetic echo trains by an echolocating bottlenose dolphin

Bottlenose dolphins (Tursiops truncatus) have an acute ability to use target echoes to judge attributes such as size, shape, and material composition. Most target recognition studies have focused on features associated with individual echoes as opposed to information conveyed across echo sequences (feature envelope of the multi-echo train). One feature of aspect-dependent targets is an amplitude modulation (AM) across the return echoes in the echo train created by relative movement of the target and dolphin. The current study examined whether dolphins could discriminate targets with different AM envelopes. "Electronic echoes" triggered by a dolphin's outgoing echolocation clicks were manipulated to create sinusoidal envelopes with varying AM rate and depth. Echo trains were equated for energy, requiring the dolphin to extract and retain information from multiple echoes in order to detect and report the presence of AM. The dolphin discriminated amplitude-modulated echo trains from those that were not modulated. AM depth thresholds were approximately 0.8 dB, similar to other published amplitude limens. Decreasing the rate of modulation from approximately 16 to 2 cycles per second did not affect the dolphin's AM depth sensitivity. The results support multiple-echo processing in bottlenose dolphin echolocation. This capability provides additional theoretical justification for exploring synthetic aperture sonar concepts in models of animal echolocation that potentially support theories postulating formation of images as an ultimate means for target identification.     

The biosonar field around an Atlantic bottlenose dolphin (Tursiops truncatus)

The use of remote autonomous passive acoustic recorders (PAR) to determine the distribution of dolphins at a given locations has become very popular. Some investigators are using echolocation clicks to gather information on the presence of dolphins and to identify species. However, in all of these cases, the PAR probably recorded mainly off-axis clicks, even some from behind the animals. Yet there is a very poor understanding of the beam pattern and the click waveform and spectrum from different azimuths around the animal's body. The beam pattern completely around an echo locating dolphin was measured at 16 different but equally spaced angles in the horizontal plane using an 8-hydrophone array in sequence. Eight channels of data were digitized simultaneously at a sampling rate of 500 kHz. The resulting beam patterns in both planes showed a continuous drop off in sound pressure with azimuth around the animal and reached levels below -50 dB relative to the signal recorded on the beam axis. The signals began to break up into two components at angles greater than ± 45° in the horizontal plane. The center frequency dropped off from its maximum at 0° in a non-uniform matter.     

Bio-inspired wideband sonar signals based on observations of the bottlenose dolphin (Tursiops truncatus)

This paper uses advanced time-frequency signal analysis techniques to generate new models for bio-inspired sonar signals. The inspiration comes from the analysis of bottlenose dolphin clicks. These pulses are very short duration, between 50 and 80 micros, but for certain examples we can delineate a double down-chirp structure using fractional Fourier methods. The majority of clicks have energy distributed between two main frequency bands with the higher frequencies delayed in time by 5-20 micros. Signal syntheses using a multiple chirp model based on these observations are able to reproduce much of the spectral variation seen in earlier studies on natural dolphin echolocation pulses. Six synthetic signals are generated and used to drive the dolphin based sonar (DBS) developed through the Biosonar Program office at the SPAWAR Systems Center, San Diego, CA. Analyses of the detailed echo structure for these pulses ensonifying two solid copper spherical targets indicate differences in discriminatory potential between the signals. It is suggested that target discrimination could be improved through the transmission of a signal packet in which the chirp structure is varied between pulses. Evidence that dolphins may use such a strategy themselves comes from observations of variations in the transmissions of dolphins carrying out target detection and identification tasks.     

Estimated communication range and energetic cost of bottlenose dolphin whistles in a tropical habitat

Bottlenose dolphins (Tursiops sp.) depend on frequency-modulated whistles for many aspects of their social behavior, including group cohesion and recognition of familiar individuals. Vocalization amplitude and frequency influences communication range and may be shaped by many ecological and physiological factors including energetic costs. Here, a calibrated GPS-synchronized hydrophone array was used to record the whistles of bottlenose dolphins in a tropical shallow-water environment with high ambient noise levels. Acoustic localization techniques were used to estimate the source levels and energy content of individual whistles. Bottlenose dolphins produced whistles with mean source levels of 146.7 ± 6.2 dB re. 1 μPa(RMS). These were lower than source levels estimated for a population inhabiting the quieter Moray Firth, indicating that dolphins do not necessarily compensate for the high noise levels found in noisy tropical habitats by increasing their source level. Combined with measured transmission loss and noise levels, these source levels provided estimated median communication ranges of 750 m and maximum communication ranges up to 5740 m. Whistles contained less than 17 mJ of acoustic energy, showing that the energetic cost of whistling is small compared to the high metabolic rate of these aquatic mammals, and unlikely to limit the vocal activity of toothed whales.     

Audibility thresholds for a pure tone in the presence of a pulse jam in the bottlenose dolphin

The effect of a pulse jam on the audibility of pure tones in the bottlenose dolphin (Tursiops truncatus) is investigated. The pulse jam consists of a sequence of pairs of identical short pulses with a pulse spacing of 50 µs in each pair and with a pair repetition rate of 300 s^?1. The test signal is represented by pure tones in the frequency range 20–100 kHz. The audibility thresholds for the test signals are measured at 10-kHz steps, both in the presence of the pulse jam and in its absence, on the basis of the conditioned-reflex method with food reinforcement. The resulting dependence of the threshold shifts (TS) due to the pulse jam on the frequency of the test signal has a complex form. This dependence can be separated into three components: (1) the oscillations of the TS curve that correlate with the extrema of the spectral density of the jam, so that the peaks and dips of the TS curve correspond to the maxima and minima of the spectral density, respectively; (2) the component monotonically decreasing as the frequency grows up to 80 kHz, which distinguishes the TS curve under consideration from the rising curve obtained for masking by random noise; and (3) the frequency-independent component of the TS curve. The following auditory features are associated with these components: component 1 determines the timbre of the pulse jam; component 2 is presumably related to the pitch corresponding to the frequency 1/τ; and component 3 exhibits a rather strong auditory feature of random noise due to the random neural activity caused by the pulse jam in the whole auditory filter band.